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Background   zic  genes are members of the  gli/glis/nkl/zic  super-family of C2H2 zinc finger (ZF) transcription factors. Homologs of the  zic  family have been implicated in patterning neural and mesodermal tissues in bilaterians. Prior to this study, the origin of the metazoan  zic  gene family was unknown and expression of  zic  gene homologs during the development of early branching metazoans had not been investigated.    Results  Phylogenetic analyses of novel  zic  candidate genes identified a definitive  zic  homolog in the placozoan  Trichoplax adhaerens , two  gli/glis/nkl- like genes in the ctenophore  Mnemiopsis leidyi , confirmed the presence of three  gli/glis/nkl -like genes in Porifera, and confirmed the five previously identified  zic  genes in the cnidarian  Nematostella vectensis . In the cnidarian  N. vectensis ,  zic  homologs are expressed in ectoderm and the gastrodermis (a bifunctional endomesoderm), in presumptive and developing tentacles, and in oral and sensory apical tuft ectoderm. The  Capitella teleta zic  homolog ( Ct-zic ) is detectable in a subset of the developing nervous system, the foregut, and the mesoderm associated with the segmentally repeated chaetae. Lastly, expression of  gli  and  glis  homologs in  Mnemiopsis .  leidyi  is detected exclusively in neural cells in floor of the apical organ.    Conclusions  Based on our analyses, we propose that the  zic  gene family arose in the common ancestor of the Placozoa, Cnidaria and Bilateria from a  gli/glis/nkl -like gene and that both ZOC and ZF-NC domains evolved prior to cnidarian-bilaterian divergence. We also conclude that  zic  expression in neural ectoderm and developing neurons is pervasive throughout the Metazoa and likely evolved from neural expression of an ancestral  gli/glis/nkl/zic  gene.  zic  expression in bilaterian mesoderm may be related to the expression in the gastrodermis of a cnidarian-bilaterian common ancestor.

Expression and phylogenetic analysis of the zic gene family in the evolution and development of metazoans